Editing Vestibulo-ocular reflex (section)

=== Oculomotor integrator ===
In addition to these direct pathways, which drive the velocity of eye rotation, there is an indirect pathway that builds up the position signal needed to prevent the eye from rolling back to center when the head stops moving. This pathway is particularly important when the head is moving slowly because here position signals dominate over velocity signals. David A. Robinson discovered that the eye muscles require this dual velocity-position drive, and also proposed that it must arise in the brain by mathematically integrating the velocity signal and then sending the resulting position signal to the motoneurons. Robinson was correct: the 'neural integrator' for horizontal eye position was found in the nucleus prepositus hypoglossi<ref name="Cannon1987">{{cite journal |vauthors=Cannon SC, Robinson DA |date=May 1987 |title=Loss of the neural integrator of the oculomotor system from brain stem lesions in monkey |journal=Journal of Neurophysiology |volume=57 |issue=5 |pages=1383–409 |doi=10.1152/jn.1987.57.5.1383 |pmid=3585473}}</ref> in the medulla, and the neural integrator for vertical and torsional eye positions was found in the [[interstitial nucleus of Cajal]]<ref name="Crawford1991#2">{{cite journal |vauthors=Crawford JD, Cadera W, Vilis T |date=June 1991 |title=Generation of torsional and vertical eye position signals by the interstitial nucleus of Cajal |journal=Science |volume=252 |issue=5012 |pages=1551–3 |bibcode=1991Sci...252.1551C |doi=10.1126/science.2047862 |pmid=2047862 |s2cid=15724175}}</ref> in the midbrain. The same neural integrators also generate eye position for other conjugate eye movements such as [[saccade]]s and [[smooth pursuit]].

The integrator is [[Leaky integrator|leaky]], with a characteristic leaking time of 20 s. For example, when the subject is sitting still and focusing on an object, and suddenly the light is turned off, the eyes would return to their neutral position in around 40 seconds even as the subject is attempting to keep the focus.<ref name="Robinson 33–45">{{Cite journal |last=Robinson |first=D A |date=March 1989 |title=Integrating with Neurons |url=https://www.annualreviews.org/doi/10.1146/annurev.ne.12.030189.000341 |journal=Annual Review of Neuroscience |language=en |volume=12 |issue=1 |pages=33–45 |doi=10.1146/annurev.ne.12.030189.000341 |pmid=2648952 |issn=0147-006X}}</ref><ref>{{Cite journal |last1=Sanchez |first1=Katherine |last2=Rowe |first2=Fiona J. |date=March 2018 |title=Role of neural integrators in oculomotor systems: a systematic narrative literature review |url=https://onlinelibrary.wiley.com/doi/10.1111/aos.13307 |journal=Acta Ophthalmologica |language=en |volume=96 |issue=2 |pages=e111–e118 |doi=10.1111/aos.13307 |pmid=27874249 |issn=1755-375X}}</ref>